Jennifer L. Kelley

Predation Risk Shapes Social Networks in Fission-Fusion Populations

Predation risk is often associated with group formation in prey, but recent advances in methods for analysing the social structure of animal societies make it possible to quantify the effects of risk on the complex dynamics of spatial and temporal organisation. In this paper we use social network analysis to investigate the impact of variation in predation risk on the social structure of guppy shoals and the frequency and duration of shoal splitting (fission) and merging (fusion) events. Our analyses revealed that variation in the level of predation risk was associated with divergent social dynamics, with fish in high-risk populations displaying a greater number of associations with overall greater strength and connectedness than those from low-risk sites. Temporal patterns of organisation also differed according to predation risk, with fission events more likely to occur over two short time periods (5 minutes and 20 minutes) in low-predation fish and over longer time scales (>1.5 hours) in high-predation fish. Our findings suggest that predation risk influences the fine-scale social structure of prey populations and that the temporal aspects of organisation play a key role in defining social systems.

Condition-dependent expression of pre- and postcopulatory sexual traits in guppies

Female choice can impose persistent directional selection on male sexually selected traits, yet such traits often exhibit high levels of phenotypic variation. One explanation for this paradox is that if sexually selected traits are costly, only the fittest males are able to acquire and allocate the resources required for their expression. Furthermore, because male condition is dependent on resource allocation, condition dependence in sexual traits is expected to underlie trade-offs between reproduction and other life-history functions. In this study we test these ideas by experimentally manipulating diet quality (carotenoid levels) and quantity in the guppy (Poecilia reticulata), a livebearing freshwater fish that is an important model for understanding relationships between pre- and post-copulatory sexually selected traits. Specifically, we test for condition dependence in the expression of pre- and postcopulatory sexual traits (behavior, ornamentation, sperm traits) and determine whether diet manipulation mediates relationships among these traits. Consistent with prior work we found a significant effect of diet quantity on the expression of both pre- and postcopulatory male traits; diet-restricted males performed fewer sexual behaviors and exhibited significant reductions in color ornamentation, sperm quality, sperm number, and sperm length than those fed ad libitum. However, contrary to our expectations, we found no significant effect of carotenoid manipulation on the expression of any of these traits, and no evidence for a trade-off in resource allocation between pre- and postcopulatory episodes of sexual selection. Our results further underscore the sensitivity of behavioral, ornamental, and ejaculate traits to dietary stress, and highlight the important role of condition dependence in maintaining the high variability in male sexual traits.

Colour change and assortment in the western rainbowfish

Grouping behaviour is widespread across the animal kingdom, and is known to reduce an individuals risk of predation, for example through predator confusion. Theory predicts that individuals that are different in appearance to the rest of the group are at a greater risk of predation because they are more conspicuous to predators (the ‘oddity’ effect). Thus, animals should choose group mates that are the most similar in appearance to themselves. Another common antipredator tactic is crypsis (camouflage). Fishes are capable of changing colour to match their visual background, but few studies have examined how this might influence shoaling decisions, particularly in the context of the oddity effect. We induced colour pattern changes in a colourful species of freshwater fish, the western rainbowfish, Melanotaenia australis, by maintaining fish in dark and pale aquaria for 2 weeks. Analysis of the proportion of black body pigmentation confirmed that rainbowfish in dark environments developed darker colour patterns than those held in pale environments. We then conducted behavioural observations to determine whether fish subsequently based their shoaling decisions on body coloration. We found that rainbowfish preferred to shoal with similar individuals; fish that had been held in dark aquaria preferred to shoal with other dark fish and fish from pale aquaria preferred other pale fish. Our findings are consistent with the predictions of the oddity effect and demonstrate how morphological colour pattern changes and behavioural decisions interact to mediate antipredator tactics in fish.

Implications of multiple mating for offspring relatedness and shoaling behaviour in juvenile guppies

Polyandry (female multiple mating) can confer important benefits to females, but few studies have considered its potential costs. One such cost may arise through differences in the relatedness of offspring born to females with different mating histories; offspring born to monandrous females are always full siblings, while those produced by polyandrous females may be full or half siblings. These differences may have important consequences for social interactions among offspring. We used artificial insemination in the guppy (Poecilia reticulata), a promiscuous live-bearing fish, to evaluate shoaling behaviour in polyandrous and monandrous broods. We combined this information with known parentage data for the polyandrous broods to determine whether sibling relatedness influenced offspring shoaling behaviour. While we detected no effect of mating treatment (polyandry/monandry) on shoaling behaviour, we found that pairs of full siblings spent significantly more time shoaling (and in close proximity) than pairs of half siblings. This latter finding confirms the ability of newborn guppies to distinguish brood mates on the basis of kinship, but also suggests an important and hitherto unrealized potential cost of polyandry: a reduction in within-brood relatedness with potentially important implications for offspring social behaviour.

Linking stream ecology with morphological variability in a native freshwater fish from semi‐arid Australia

Environmental variation is a potent force affecting phenotypic expression. While freshwater fishes have provided a compelling example of the link between the environment and phenotypic diversity, few studies have been conducted with arid-zone fishes, particularly those that occur in geographically isolated regions where species typically inhabit intermittent and ephemeral creeks. We investigated morphological variation of a freshwater fish (the western rainbowfish, Melanotaenia australis) inhabiting creeks in the Pilbara region of northwest Australia to determine whether body shape variation correlated with local environmental characteristics, including water velocity, habitat complexity, predator presence, and food availability. We expected that the geographic isolation of creeks within this arid region would result in habitat-specific morphological specializations. We used landmark-based geometric morphometrics to quantify the level of morphological variability in fish captured from 14 locations within three distinct subcatchments of a major river system. Western rainbowfish exhibited a range of morphologies, with variation in body depth accounting for a significant proportion (>42%) of the total variance in shape. Sexual dimorphism was also apparent, with males displaying deeper bodies than females. While the measured local habitat characteristics explained little of the observed morphological variation, fish displayed significant morphological differentiation at the level of the subcatchment. Local adaptation may partly explain the geographic patterns of body shape variation, but fine-scale genetic studies are required to disentangle the effects of genetic differentiation from environmentally determined phenotypic plasticity in body shape. Developing a better understanding of environment–phenotype relationships in species from arid regions will provide important insights into ecological and evolutionary processes in these unique and understudied habitats.

Male sperm storage compromises sperm motility in guppies

Sperm senescence can have important evolutionary implications due to its deleterious effects on sperm quality and offspring performance. Consequently, it has been argued that polyandry (female multiple mating) may facilitate the selection of younger, and therefore competitively superior, sperm when ejaculates from multiple males compete for fertilization. Surprisingly, however, unequivocal evidence that sperm ageing influences traits that underlie sperm competitiveness is lacking. Here, we used a paired experimental design that compares sperm quality between ‘old’ and ‘young’ ejaculates from individual male guppies (Poecilia reticulata). We show that older sperm exhibit significant reductions in sperm velocity compared with younger sperm from the same males. We found no evidence that the brightness of the males orange (carotenoid) spots, which are thought to signal resistance to oxidative stress (and thus age-related declines in sperm fitness), signals a males ability to withstand the deleterious effects of sperm ageing. Instead, polyandry may be a more effective strategy for females to minimize the likelihood of being fertilized by aged sperm.

Conflict between background matching and social signalling in a colour-changing freshwater fish.

The ability to change coloration allows animals to modify their patterning to suit a specific function. Many freshwater fishes, for example, can appear cryptic by altering the dispersion of melanin pigment in the skin to match the visual background. However, melanin-based pigments are also used to signal dominance among competing males; thus colour change for background matching may conflict with colour change for social status signalling. We used a colour-changing freshwater fish to investigate whether colour change for background matching influenced aggressive interactions between rival males. Subordinate males that had recently darkened their skin for background matching received heightened aggression from dominant males, relative to males whose coloration had not changed. We then determined whether the social status of a rival male, the focal males previous social status, and his previous skin coloration, affected a males ability to change colour for background matching. Social status influenced skin darkening in the first social encounter, with dominant males darkening more than subordinate males, but there was no effect of social status on colour change in the second social encounter. We also found that the extent of skin colour change (by both dominant and subordinate males) was dependent on previous skin coloration, with dark males displaying a smaller change in coloration than pale males. Our findings suggest that skin darkening for background matching imposes a significant social cost on subordinate males in terms of increased aggression. We also suggest that the use of melanin-based signals during social encounters can impede subsequent changes in skin coloration for other functions, such as skin darkening for background matching.

Morphological plasticity in a native freshwater fish from semiarid Australia in response to variable water flows

Abstract In fishes, alterations to the natural flow regime are associated with divergence in body shape morphology compared with individuals from unaltered habitats. However, it is unclear whether this morphological divergence is attributable to evolutionary responses to modified flows, or is a result of phenotypic plasticity. Fishes inhabiting arid regions are ideal candidates for studying morphological plasticity as they are frequently exposed to extreme natural hydrological variability. We examined the effect of early exposure to flows on the development of body shape morphology in the western rainbowfish (Melanotaenia australis), a freshwater fish that is native to semiarid northwest Australia. Wild fish were collected from a region (the Hamersley Ranges) where fish in some habitats are subject to altered water flows due to mining activity. The offspring of wild‐caught fish were reared in replicated fast‐flow or slow‐flow channels, and geometric morphometric analyses were used to evaluate variation in fish body shape following 3, 6, 9, and 12 months of exposure. Water flows influenced fish morphology after 6 and 9 months of flow exposure, with fish in fast‐flow environments displaying a more robust body shape than those in slow‐flow habitats. No effect of flow exposure was observed at 3 and 12 months. Fishes also showed significant morphological variation within flow treatments, perhaps due to subtle differences in water flow among the replicate channels. Our findings suggest that early exposure to water flows can induce shifts in body shape morphology in arid zone freshwater fishes. Morphological plasticity may act to buffer arid zone populations from the impacts of anthropogenic activities, but further studies are required to link body shape plasticity with behavioral performance in habitats with modified flows.

The Biological Mechanisms and Behavioral Functions of Opsin-Based Light Detection by the Skin

Light detection not only forms the basis of vision (via visual retinal photoreceptors), but can also occur in other parts of the body, including many non-rod/non-cone ocular cells, the pineal complex, the deep brain, and the skin. Indeed, many of the photopigments (an opsin linked to a light-sensitive 11-cis retinal chromophore) that mediate color vision in the eyes of vertebrates are also present in the skin of animals such as reptiles, amphibians, crustaceans and fishes (with related photoreceptive molecules present in cephalopods), providing a localized mechanism for light detection across the surface of the body. This form of non-visual photosensitivity may be particularly important for animals that can change their coloration by altering the dispersion of pigments within the chromatophores (pigment containing cells) of the skin. Thus, skin coloration may be directly color matched or “tuned” to both the luminance and spectral properties of the local background environment, thereby facilitating behavioral functions such as camouflage, thermoregulation, and social signaling. This review examines the diversity and sensitivity of opsin-based photopigments present in the skin and considers their putative functional roles in mediating animal behavior. Furthermore, it discusses the potential underlying biochemical and molecular pathways that link shifts in environmental light to both photopigment expression and chromatophore photoresponses. Although photoreception that occurs independently of image formation remains poorly understood, this review highlights the important role of non-visual light detection in facilitating the multiple functions of animal coloration.

Group size and associative learning in the Australian magpie (Cracticus tibicen dorsalis)

Group living can present unique challenges that may require individuals to possess cognitive skills, such as the ability to recognise group members and maintain relationships with specific individuals. These skills may be particularly important for animals that live in large groups, because the intricacies of social life may become more complex when more individuals are involved. Previous research has found that species with regular social interactions tend to show elevated cognitive performance relative to those that rarely interact, yet intraspecific variation in performance among individuals in social groups of varying size is rarely explored. We investigated the relationship between the ability to solve an associative learning task and group size among individuals of a free-living, social bird, the Australian magpie (Cracticus tibicen dorsalis). Individuals varied in their likelihood of interacting with and solving the task. Individuals from larger groups were more likely to approach the associative learning task, suggesting that group size influences individual propensity to attempt a novel task. However, group size did not influence the likelihood that individuals solved the task. Rather, age had an important effect; adults were more likely to solve the association task than juveniles. Our finding that free-living individuals occurring in large social groups were more likely to interact with a novel task suggests that group size may affect differences in performance at a cognitive task within a species.